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464-8602ĦƢƟÒÎĉĊÂƪÁǋƓMichitaka Notaguchi
Systemic long-distance transport of RNAs in plants
Key words: Grafting, Mobile RNA, Phloem, RNA-Seq analysis, Systemic signaling
Graduate School of Bioagricultural Sciences, Nagoya University,
Institute of Transformative Bio-Molecules, Nagoya University, JST PRESTO
Furo-cho, Chikusa-ku, Nagoya 464-8602, JAPAN
DOI: 10.24480/bsj-review.8b9.00119

 '(åƋ.ƎƆ/ȽRNA Bȗģë*( #*ǫA?( >Ȼ“RNA world Ǵ”, Joyce 2002ȼ
!.ęȽDNA-]iPǺ.Ɣ3ƨǞ#Ɗä9ȽRNA ³ø/ƎƆ.ƹǏn~[Y.ŠčB,ȽDNA ǣǢBȣö>nzGs*(ȽȗģëƗƊ>Ȫ.v_[Xx*(Ƚ{q\u.Ǫ÷Ğ*(©,+ȽƎƆ-ğǤÏū,ìȯ)Ũ-ŪǐBŜ#( >Ȼâ 1Aȼ
ţƆǦƀBē(>*ȽţƆȎÀ.ȑƩ)Ɖě# RNA ƊǸ.'*(ȽƴƲB# RNA .ȢǾȭA

B

長距離移行する植物のRNA

防御応答 などのRNA 分子種

全身性の応答様々な環境要因

tRNA
rRNA
mRNA

篩管細胞

miRNA

篩部伴細胞

siRNA

栄養分配 などの 環境応答( &)602%:57- (

Ũ, RNA ³øƪȻAȼȽƴȘƹǏ<ÈĔǺȍǀB(ƴƲƹǏ-Ƭ6Ȇ5?>ƴƲ.sYl~ȻǻơÆȼ-&(ȽåȘ.īȢƀ:å Ș.Š-Ó(­ȁ3*ȢǾȭƨǞ>ƨǞ«- ( RNA ³øãŐ-ĵ'Ǝƌ¡ƐBƗļȽ¨}o|)ǁ¹.*?#ƗƎīȢBŜ# #% &!")
 $'"(       ƨǞƢ<?>Ȼâ 1Bȼ
ţƆ.ƴƲ/ȽƎ#ƹǏ<,=Ƚ¬ÐīƏƆȽ,A%Şȸ³B­ȁ-Ȑ2©Bĵ'ňȽƴƲ/­ȁ.ßüB',Ƽǆ*(Ƚ¨. >Ș)ħÌ#ģëB¶.Ș-ȓ>WQc|ȓƽǿ*(©*9ȇČŊ<-,='' >ƴƲ/ȋȒƼǆ
3*ƹǏ³À>ȑƩ)şBô ȽȘ.L|Nfz*ƹǏǺ-;&(ŧī?#ƹǏ*,>!
.#8Ƚş-¿Ő?>SguģëȂ±ǊǮ?>*/³À#ƴƲƹǏ)/Ǽ=, !?-9ȥA<ȽƴƲ.-/Şȸ³Ƚp|w,+.³øÀÐƆ.-ȽRNAȽ]iPǺ*&#ƎȺ³øùä>*Ƚ1990 ČȨ.ƴƲŻ³ŚBȋ(ġȌ-Ŋ<*,&#ȻLough and Lucas 2006, Notaguchi and Okamoto 2015ȼĚ&(ȽRNA :]iPǺ×=.ƹǏ<ƴƲƹǏ3*Ƭ6Ȇ5?>* *) >ƴƲƹǏ.ȫ-/ƴȘƹǏä=Ƚ]iPǺ: RNA.Ƭ6Ȇ6/ƴȘƹǏ<ÈĔǺȍǀB(ǞA?>*ǌ<?( >ƴƲBȐ0?> RNA³ø.ƪȴ/òŨ) =ȽmiRNA, siRNA, tRNA, long noncoding RNA, mRNA ,+Ƚȋċ.ƹǏ)©³øƪ.òȽ!.ŧī/ƕ,>9..ȽƴƲ-9ǥ'>ŬWRUBƐ #SguGb, RNA ǩŚĮǟ-;=ȽƴƲ. RNA ³ø-' (¿Ĵƙ,ģëě<?>ŋ*,=Ƚ
ęƚ.ƣƭ.ȵȀǥȆ5?>Ŗƫ)/ȽƴƲB(ţƆ.­ȁBȢǾȭƨǞ> RNA ³
ø.%Ƚƃƀ?(ƣƭȎ8<?(# small RNA * mRNA -' (ŦǴ>


%:<9=57  ȇČ.sGP~E}G:ŬWPJUBƐ #ǩŚ-;&(ȽǗB´ņ(ě<?>ƴƲ
Ż.-/òŨ, small RNA Ô5?>*Ŋ<*,&#ȻMolnar et al. 2010, Buhtz et al. 2010ȼ

miRNA : siRNA * &# small RNA /ƴƲB(­ȁ3ȢǾȭȄȊ?ȽȗøUG}WQȽ
ƍïĠƱȽŞȸș³Ƚ!(ƗƎB¹Ĝ>*ǌ<?( >ȻKehr and Buhtz 2007, Mlotshwa et al.
2002ȼ siRNA -' (/Ƚ­ȁƨǞ> RNA .)ŪǐŎ9ǴŊ?( >siRNA .­ȁƨǞĢ' (/ siRNA ƏƎ-ȥA> Dicer .ðƕBƐ #ĸœþȹ-;&(Ŋƥ-Ʀ?(=ȽsiRNA.­ȁƨǞ-;&(Ȃ±}o|.ȗøUG}WQȻTranscriptional gene silencing, TGSȼ*Ȃ±ę- mRNA B³ǩ>*)Ǽ>Ȃ±ęȗøUG}WQȻPost-transcriptional gene silencing,
PTGSȼ.ňBđǼ*Ƣ<?>?<.ȗøUG}WQŪŧ/HF|Y,+.Ɩ
ÿ-<?#ȪȽ­ȁƙ-ĲİĢBƗļ>)Ȯċ-ȝǤ) >ȻMolnar et al., 2010ȼ
TGS -'(ƺ>*ȽTGS / siRNA .©-;&(ȽRNA-directed DNA methylation (RdDM)B# DNAv^|ÀȽ > /P~s^.ŹÀǼȽƾŜ*(ũƙ?#ȗø.Ȃ±į¹?>
Bai et al. (2011).ĸœþȹ)/Ƚ2 ƪȴ.ţƆȽM{lzwVGPHF|Y. 35S n~w]
-ƒř> siRNA BĔī>ţƆ* 35S n~w]) GFP ȗøBƗƊ>ţƆȻCaMV
35Sp::GFPȼƐ <?ȽǍ.Ȥ)ĸœBǞ*ȽCaMV 35Sp::GFP ţƆ-> GFP ȗø.
ƗƊį¹Ǽ>*Ʀ?#<-ȽsiRNA ƏƎ),Dicer .') > DCL3 .ðƕ

BƐ #ĸœþȹ)/ȽȞƎç)ǲ8<?> TGS  dcl3 ðƕǎŌ)/đǼ?, *<ȽsiRNA ­ȁƨǞ>þ) =ȽTGS / siRNA .ȄȊ«)đǼ?>*ƦÝ?# #% &!")
 $'"(       ȻMelnyk et al. 2011ȼ
RdDM ƽǿ-' (9ƌǩż5=ȽDCL3  24 îê. siRNA BƏƎ>
*ȻQi et al. 2005ȼ
ȽRdDM / 24 îê. siRNA -;=đǼ?>*Ŋ<*,&#ȻLaw andJacobsen 2010ȼ
Lewsey et al. (2016) /­ȁƨǞĢ siRNA -;> DNA v^|À-' (SguGb-ǩŚȽsiRNA -;> RdDM / DOMAINS REARRANGED METHYLTRANSFERASE 1 (DRM1) *DRM2 -£ù(Ǽ>*BŊ<-#BÐA >*ȽTGS /5 DCL3 -;> 24 îê. siRNA .ƏƎ-ö5=ȽŬ-ƏƎ?# siRNA .­ȁƨǞȽƨǞ«). DRMs *.ǣÐBĔī
;1 RdDM .Ɨ¾*ǂȽŎƻƙ-ũƙȗø.ȗøUG}WQƥƮ?>* .Ɗƈ.ƌǩ) >,ȽDNA v^|ÀƈĨ.ðÀ/ȽW~GecZc.ƕ,>JT]GnȤ)ĸœ
#ìÐ:ȻLewsey et al. 2016ȼ
ȽH{Ƨ:cYƧ.ƕ,>ƪȤ)ĸœ#ìÐ-9Ť²?(=ȻAvramidou et al. 2015, Kasai et al. 2016, Wu et al. 2013ȼȽąřƙ-/þƐ¡Ɔ- (9ȽĸœB(JkXIf`F_P,ðÀBǳĆ>*)ŐƐĔǺB>* &#ĮǟƎ5?>ÏǐĢ> ňȽmiRNA .ƨǞ-ȥ>Ɨǥ/ȽǤŞȸƸ.') >{ȻPȼ3.ĠƱŨĐ.ƣƭ-/5>{/ȽATPȽşțȽ{ǑǺ-Ô5?>ğȱªƸ) =ȽƂŪ{țȻPiȼ*(Š<Ë=Ȇ5?>ȻSchachtman et al. 1998ȼ
Pi /ȽǃƲŗBȐ0?>Ȫ.Ǥ,ĔĨ) =ȻBieleski 1973ȼȽ
Pi .ūŋ-/ȽūƈĨBǡ#8 Pi .ÕÊ¼/Ⱥ5>ȻDrew and Saker 1984ȼ
Pi .ūƈĨ-?#ţƆ/ȽmiRNA .ƪ) > miR399 BȺƗƊȽmiR399 .ũƙ*,> PHO2 ȗø.ƗƊ
Bį¹>PHO2 ȗø/yjO^ƾÐȚƸBTbȽmiR399 -;> PHO2 ȗø.ƗƊį¹/Ƚ!. ź- ( Pi azYr]ȗø.ƗƊŉBđǼȽ!.ƾŜȽǚ->Pi .ǜƬ¥Ȏ?>ȻBari et al. 2006, Fujii et al. 2005ȼ
.;-ȽmiR399/PHO2 WQc|ȓƽǿ-;&(Ƚ{.­ȁƙ,ș³ǵŃ?( >!.ę.ƣƭ)Ƚ.vMdZu/Ũ,ţƆď¦ù?( >*9Ɗä)/Ƣ<?( >ȻBranscheid et al. 2010, Zhang et al. 2016aȼ{ñ.ŞȸƸ-' (9Ƚ­ȁƙ,ș³.ǵŃ.#8-ȴ#Ūŧ©ìÐǥ'&(=ȽƤțî.ūŋ-9 miR395 *  miRNA .ƗƊŉǲ8<?ȽƾŜ*(ƤțazYr]ȗø.ƗƊŉ>*ëÖ?( >ȻBuhtz et al. 2010, Kawashima et al. 2011ȼ
-XxNGw)/ miR172 * miR156 .ƨǞĢëÖ?(=Ƚ!?</Ǖī:íǗĔī.]GtQB¹Ĝ(>ÏǐĢƦÝ?( >ȻMartin et al. 2009; Kasai et al. 2010; Bhogale et al. 2014ȼ
ȽmiRNA /ƹǏǒĘƙ-©ìÐǔƙ) >ȻSchwab et al. 2006ȼȽǭ.¢- >;-ȽȘ. miRNA-' (/¨}o|.­ȁĢ.WQc|ȓȻȢǾȭWQc|ȓ*Ù0?>ȼ-©*ƦÝ
?( >,Ƚ30 90 îê* si/miRNA ;=/Ȣsmall noncoding RNA -' (9 in vitro )+C,ŹĢBĵ'ǵ4<?#* =Ƚ]iPǺǊǮ.ȧÿŹĢBƦ*ëÖ?( >ȻZhang et al. 2009ȼ
.;-ȽţƆ.¨.-/Ũ, small RNA ȢǾȭƨǞȽ³øƪ*-ƕ,>ŨĐ)ȽƨǞ«.Ƽǆ->ũƙ³ø.ƗƊ}o|BǵƳ( >
%:<9=57  mRNA .ƴƲB#ȢǾȭȄȊ/Ƚ?5)-ǣł.ţƆƪBƐ (ƣƭ?((=ȽȄȊ #% &!")
 $'"(       ?> mRNA .Ñý:!.ȄȊŪŧ-ȥ>Ƣǥ9Ȭ5&(( >ǓØż *-Ƚǣł.ţƆ
)ȢǾȭƨǞ> mRNA BÑý>*ȽƇý.ȗølDt{. mRNA ǵ4#ǣł.ƪ)®ȋ
(ǥ'>*³=ȽmRNA .ȢǾȭƨǞ/¦ù?#ƊǸ) >*Ĺă?( >ȻNotaguchi 2015 -(ŦǴȼ<-ȽƴƲ-/ RNA ƾÐ]iPǺǥ'>*ȻBalachandran
et al. 1997, Xoconostle-Cázares et al. 1999, Aoki et al. 2002, Gómez et al. 2005ȼ
Ƚ!?<.]iPǺ/ũƙ
mRNA .Ƈý.șµ-Ą(Ƈƕƙ-ƾÐȽȢǾȭƨǞ.#8. RNA]iPǺǣÐBĔī
>*ƦÝ?( >ȻHam et al. 2009, Cho et al. 2012ȼ
5#ȽƴƲ-/!9!9 RNA ³ǩȚƸ.ŹĢŤ²?, *9Ƣ<?ȻSasaki et al. 1998, Doering-Saad et al. 2002ȼȽţƆ-ħŝ>ţƆ
HF|YȻò/ RNA HF|Yȼ/!<ā.ţƆŐ>ƴƲȄȊWY`uB·Ɛ( >* Ĺă9ǔƙ-,?( >ȻNelson and Citovsky 2015ȼ
/ȽţƆ/ <. mRNA B
ȢǾȭȄȊ>ŪŧBƉěǃĵ(=ȽȄȊ?> mRNA -/ <.ƎƆûƙĥǉ >*BĹă >Ƚ?5)-ƴƲBƨǞ> mRNA .ƣƭ/Ƶ¼ƙ-ǞA?( >9..Ƚ
!.Ė»-' (/Ů+Ŋ<-?( , Ǳƹ-' (/ȽŖƫñ-9¶.ŦǴǶńBÉƅ
?# ȻKehr and Buhtz 2007, Harada 2010, Spiegelman et al. 2013, Notaguchi 2015ȼ Ŏȇ-,=ŬWRUBƐ #SguGb,ǩŚǞA?Ƚĸœ:ĂƎ* &#ƪȴ.ţƆĸÐ#ìȯ)/Ƚ!?<.ţƆ.Ȥ)łƘ<łÂ. mRNA ĸÐȘBǽ(ȢǾȭȄȊ?>*ëÖ?( >ȻKim et al. 2014, Notaguchi et al. 2015, Thieme et al. 2015, Yang et al. 2015,
â 2 A, BȼȘ. mRNA -' (/<,>ŪǐǩŚǰ6<?ȽmRNA ȢǾȭȄȊ?#ƾŜȽȄȊ?#«.ƗƎƊǸ-ĕȰB>*ëÖ?( >ȻKim et al. 2001, Haywood et al. 2005,
Banerjee et al. 2006, Notaguchi et al. 2012, â 2Cȼ
?5).ƣƭ)/ȽȄȊ«) mRNA ]iPǺ

A

B
mRNA

RNA-Seq 解析

タバコ属
（Nb ）

遺伝子1

At

Nb

遺伝子2
遺伝子3

Nb
At

C
穂木

欠損

優性

台木

欠損

欠損

・

12

・
・

接ぎ木
部位
シロイヌナズナ
（At ）

・

8

・

6

・
・

4

・

数百〜数千 の長距離移行性mRNA
の同定

・
・
・
・
・

アクチン

( 1#$"!"402%:57- 

P >0.01 P >0.01

10・
Illumina GIIx Sequencer

側根数
14

1 cm

2
0

野生型 欠損 優性 欠損 優性
欠損 欠損

'*

ȻAȼƪȴ.ţƆBĸœ>*Ƚ­ȁƨǞĢ. mRNA /ĸœȘBǽ(Ɲĭ.ţƆ3ƨǞ>Ŭ
WRUBƐ (Ɲĭ.ţƆ.mRNA BǄǇǩŚ>*)Ƚ
ƨǞ(#mRNA .șµģëBËě)Ƚ
­ȁƨǞĢ. mRNA BÑý>*)>
ȻBȼRT-PCR -;>ƾŜ.ȉǯ
ȻCȼ­ȁƨǞĢ mRNA *(Ñý?# Aux/IAA mRNA .Ǝƌ¡Ɛ-' (ŤǬ#ĸœþȹ!?"?.þȹ.Ǳƹ/ȽÈǛǶńBÉƅ
ȻA, B: Notaguchi et al. 2015, C: Notaguchi et al. 2012ȼ #% &!")
 $'"(       

3ǊǮ?( >*BŊƦ#¢/, ȽǠƊç.ǩŚ<]iPǺ3.ǊǮ/Ǽ=>*Ĺă?( >ǛǍ<.º.ŤǬ)/ȽȢǾȭȄȊ?> mRNA ǈ.ƇĝBSgu.­ mRNA.Ƈĝ*űȃ(6>*Ƚ mRNA .Ȣ-9Ƚȗøn~w].șµw^l-9ȽmRNA!.9..șµ-9ȽƇư4Ƈĝ/ǥ'<,&#ȻNotaguchi et al. 2015ȼǓØż&#./Ƚ
ƨǞĢBƦ mRNA .-/Ƚ/8<­ȁ)ƗƊ>hHYOkQȗø9òÔ5?(#*) >!?<. mRNA  (¨.BƨǞ>ğǤĢ/ ;-ǌ<?>#8ȽƨǞĢ mRNA .-/WQc|ȓ-ȥ¤>9.*ȥ¤, 9..ňÔ5?( >.)/,
* .ǛǍ.Ĺă) >,ȽȢǾȭȄȊ?> mRNA .ƗƊȟ*ƨǞĢ.ȥ¤Bǵ4>*ȽÅƷ,ƗƊȟƨǞĢBǴŊ>Ǯ)/,ȽƨǞ-/ <.¹ĜŪŧ©*Ĺă?
#ȻNotaguchi et al. 2015ȼ
#$Ƚ¶.ƣƭQ|n.ǩŚƾŜ)/Ƚò. mRNA .ìÐ-' (ƗƊȟ*ƨǞĢ.Ȥ-ȥ¤Ģǥ²?>*9ƦÝ?(=ȽƴȘƹǏ<ƴƲƹǏ3.
mRNA .ƨǞ-' (/ȽȖıĢBĵ&#¹ĜBÌ>ƨǞ*ȽƗƊȟ-Ġ#ý}o|.žŸȽÇ%ȖıĢ. ȽȖıƙ¹ĜBÌ, ƨǞ.i] =>*ǌă?( >ȻThieme et al.
2015, Calderwood et al. 2016ȼȴ#ƴƲƹǏ3.ƎȺ³ø.žŸ/Ƚ]iPǺ.ìÐ)9ǲ8<?>* ëÖŎȇ,?(=ȻPaultre et al. 2016ȼȽę/!#ý.žŸƈŵBǌĩ(
ƾŜ*ÓÐğǤ >
 5#þȹĭŶ.Ǩƀ)9ŷĥ/ğǤ) >ƆǺ.ȢǾȭȄȊ.Ťǯ-;Ɛ <?>ĸœŶ' (ȽǓØż ëÖ,?( >Yang et al. (2015) -;>*Ƚĸœ( 11 Čƽȑ#mbH
.īœǖ*ĸœ( 1 ŏ.ĎǖBƐĥȽ!?"?.ǰŅ<ƨǞĢBƦ mRNA BÑý(6>*Ƚ11 Čǖ)/ 1 ŏǖ.ìÐ-ű4(ƨǞĢBƦ mRNA łȻƪȴłȼć,,>* .
ƾŜ.ǩȜ*(ȽīȢY`X-;&(ƨǞ> mRNA .ƪȴðÀ#* ÅƷ,ǩȜ.-Ƚĸœ* ƁĿ¡-;>ĕȰŋȤƽȑ*®-Ǚ%Ơ ##8*>ǩȜ9ī=Ʈ'!.#8Ƚ
mRNA .ƨǞĢ-' (Ŏƻƙ,ƾǶBě>-/Ƚĸœñ.ĭŶ-;>ǰȹ9ğǤ) >*ǛǍ/ǌ> mRNA .ƨǞ-' (/ȮȖıƙ,žŸ.ÏǐĢBĶ#Ƚý.¹ĜŪŧ© ( >*9
?5).ƣƭ)Ŋ<-?( >ƨǞĢBƦ mRNA .Ș³șµBȽŖř/ƨǞĢBƦ,
GFP : GUS ,+.}r]ȗø-ǝÐ>*ȽGFP : GUS mRNA -ƨǞĢ?>*þȹƙ-Ʀ?( >ȻBanerjee et al. 2009, Huang and Yu 2009, Thieme et al. 2015, Zhang et al. 2016bȼ?5)-ǣł. mRNA BĄǸ-þȹǞA?ȽmRNA . UTR Ș³,+Ƈý.șµȳèƨǞȝǤ) >*Ʀ?( >«Ȉ.;- RNA ƾÐ]iPǺƴƲ-ùä>*:Ƚ
-9ȽƨǞĢBƦ mRNA .¶.ȗølDt{Bǵ4>*Ƚğ9ƨǞĢBƦǮ)/, *,+ȻHuang and Yu 2009, Notaguchi et al. 2012, 2015ȼ
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